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Home , Azure gallinule, Gallirallus, Huahine, New Caledonian gallinule, Rail (bird), Society Islands

New of Rails (Aves: Rallidae) from an Archaeological Site on , Society Islands1

Jeremy J. Kirchman2,4 and David W. Steadman3

Abstract: We examined 50 bones previously assigned to ‘‘ new sp.’’ from the prehistoric (1,250–750 yr B.P.) Fa‘ahia archaeological site on Huahine, . Most of these specimens ðn ¼ 47Þ, representing nearly all major cranial and postcranial skeletal elements, belong to a medium-sized flightless that we name Gallirallus storrsolsoni. Three femora represent a second species of extinct rail that we name Porphyrio mcnabi. With the description of these two species of rails, the total number of extinct species of land from the Fa‘a- hia site stands at seven, consisting of two rails, two doves, two parrots, and a starling. Fa‘ahia also has yielded bones of six other species of land birds that no longer exist on Huahine but survive elsewhere in .

Fossil bones from archaeological and The Fa‘ahia site (Figure 2) was excavated paleontological sites on islands throughout by Yosihiko H. Sinoto and colleagues from Oceania have revealed extensive 1973 to 1984 in cooperation with the De´part- of birds after colonization by hu- ment de Arche´ologie, Centre Polyne´sien des mans and their commensals ( James and Ol- Sciences Humaines, (dapt). The cul- son 1991, Olson and James 1991, Steadman tural deposits at Fa‘ahia were found sub- 1995, in press, Worthy and Holdaway 2002). merged below the modern water table and On a typical island in East (Figure contained exceptionally well-preserved or- 1), 50 to 100% of the species of land birds ganic materials, including wooden adze that existed at human contact do not survive handles, tapa beaters, and parts of canoes there today (Steadman in press). Especially (Sinoto 1975, 1979). Other objects preserved prevalent among the East Polynesian extinct and recovered from Fa‘ahia include non- species are rails (Rallidae), most of which human bones representing food items of all were flightless species endemic to single is- sizes (fishes, reptiles, birds, and mammals) lands or to islands connected in the Pleisto- and a wide variety of artifacts and ornaments cene during periods of lowered sea levels. In made of bone, shell, or stone. By East Poly- this paper we describe two new species of nesian standards, Fa‘ahia is an early occupa- rails from the Fa‘ahia archaeological site on tion site with radiocarbon dates ranging from Huahine in the Society Islands. ca. 1,250 to 750 yr B.P. (¼ ca. A.D. 700 to 1200 [Sinoto 1983]). We note, however, that the chronology of human colonization in 1 Financial support came from a McGlaughlin Fel- lowship to J.J.K. from the University of Florida College East Polynesia (the eastward, of Liberal Arts and Sciences, and NSF grants EAR- including the Society Islands) has been de- 9714819 and DEB-0228682 to D.W.S. Manuscript ac- bated for decades (Sinoto 1970, Kirch 1984, cepted 20 June 2005. 1986, 2000, Spriggs and Anderson 1993, 2 Department of Zoology, University of Florida, Conte and Anderson 2003). Archaeological Gainesville, Florida 32611. 3 Florida Museum of Natural History, University of sites older than A.D. 1000 are scarce or ab- Florida, Gainesville, Florida 32611. sent in East Polynesia, although sedimento- 4 Corresponding author: e-mail: jkirchman@zoo logical and paleobotanical information .ufl.edu; phone: 352-392-1721 x232; fax: 352-392-3704. suggests that people were present in the Soci- ety Islands and Cook Islands at least several Pacific Science (2006), vol. 60, no. 2:281–297 centuries earlier than A.D. 1000 (Lepofsky et : 2006 by University of Hawai‘i Press al. 1992, 1996, Kirch and Ellison 1994, Lep- All rights reserved ofsky 1995). In spite of uncertainty in the

281 Figure 1. Oceania. Dashed line indicates the East Polynesia faunal . Rails from Huahine . Kirchman and Steadman 283

Figure 2. Society Islands, with inset of Huahine showing the location of the Fa‘ahia archaeological site.

chronology of first arrival, there is a consis- three were from tabuensis, a small tent pattern of heavy exploitation of native species that is widespread in Polynesia but birds early in the archaeologically preserved that no longer occurs on Huahine. The other cultural sequence in East Polynesia, which 50 bones were assigned to a larger, presum- typically begins at ca. A.D. 1000 (Dye and ably undescribed, flightless rail, referred to as Steadman 1990, Kirch et al. 1995, Steadman ‘‘Gallirallus new sp.’’ These 50 specimens are and Rolett 1996). In yielding bones that the basis of this paper. mainly represent extinct species, the zooarch- aeological evidence from Fa‘ahia agrees with materials and methods this pattern. Over 300 bird bones were obtained at Skeletons used for comparisons are from Fa‘ahia by screening the sediment through the American Museum of Natural History 0.25-in. (6.4-mm) mesh. Of 53 rail bones (amnh), Bernice P. Bishop Museum (bpbm), reported by Steadman and Pahlavan (1992), Delaware Museum of Natural History 284 PACIFIC SCIENCE . April 2006

(dmnh), Florida Museum of Natural History, each species were used for the PCA. Charac- University of Florida (uf), University of ters used in the PCA were chosen on the basis Michigan Museum of Zoology (ummz), Na- of availability in fossil specimens from Hua- tional Museum of Natural History, Smith- hine. Osteological terminology follows Bau- sonian Institution (usnm), University of mel and Witmer (1993). Washington Burke Museum (uwbm), and ypm Yale Peabody Museum ( ). We examined comparative osteology and these modern specimens: Porzana (Poliolim- systematics nas) cinerea, dmnh 72836, 72906; Porzana ta- buensis, uwbm 42501; Gallirallus striatus, amnh Family Rallidae 22981, usnm 85892, 559919, ypm 107205; G. Gallirallus Lafresnaye, 1841 torquatus, amnh 17715–17717, ummz 228275, 228280, usnm 290445; G. owstoni, uf 39918, We regard all ‘‘typical long-billed rails’’ 39920, 39921, usnm 561968, 611816, from Oceania as species of Gallirallus sensu 612616, 613738–613744, 614233–614235, lato, distinguishing them from the similarly 614771, 614772; G. australis greyi, uf 24326, sized (Porphyrio), (Gal- 24327; G. a. australis, ypm 102249, 110760, linula), and (Fulica), and the much 110789, 110790, 110844; G. philippensis good- smaller crakes (Porzana and Poliolimnas). This soni, UF 39854, 39855; G. p. sethsmithi, uf treatment departs slightly from the classifica- 42902, 42933–42935; G. p. philippensis, usnm tion of Olson (1973a), who provisionally re- 560651; G. p. yorki, usnm 560791; G. p. tained woodfordi of the Solomon Islands and mellori, usnm 620196; G. p. ecaudatus, uwbm poecilopterus of Fiji in the genus Nesoclopeus 42863, 42865, 42866; G.[‘‘Nesoclopeus’’] wood- but highlighted their close affinity with Gal- fordi, uf 39399, 39406, 39409, 39547, 39556, lirallus. Livezey (1998, 2003) retained the 39574; Gallinula chloropus, uf 39927; Porphyrio genera Nesoclopeus, Tricholimnas, Cabalus, and porphyrio samoensis, uf 39332, 39388, 39407, Habropteryx for some species of long-billed usnm 561547, 561549, 5461551; P. p. polioce- rails but acknowledged the difficulty of es- phalus, usnm 34212; P. p. pulverulentus, usnm tablishing generic-level relationships in this 226035, 292296, 292297; Porphyrio martinicus, group on the basis of osteology. We refer 47 uf 42417, 42419; Porphyrio alleni, uf 34172, of the from Huahine to Gallirallus 38839. We also examined these fossil speci- rather than to other genera of Oceanic rails mens: Gallirallus ripleyi humerus, uf 51402, because of the following characters. Skull: ulnae, uf 54901, 55215, carpometacarpi, uf frontals narrow, concave. Rostrum: long, 54700, 54988, femur, uf 51320, tibiotarsi, uf narrow, and shallow with elongate nares. Hu- 54732, 54985, usnm 402895 (holotype), tar- merus: fossa pneumotricipitalis deep and wide sometatarsi, uf 54761, 55223, usnm 402895 with prominent crus ventrale fossae. Ulna: (holotype); G. vekamatolu humerus, uf thin in cranial aspect with rectangular (rather 52333, ulna, uf 51734, femora, uf 52020, than rounded) margo cranialis. Pelvis: ala 52058, tibiotarsi, uf 51729, 52211, tarsometa- preacetabularis ilii broadly continuous with tarsus, uf 51991 (holotype); Porphyrio paepae crista dorsalis of synsacrum. Femur: distal femora, bpbm 165649, 166424, 166426, end of corpus femoris becomes gradually 166434, tibiotarsus, bpbm 165651, synsacrum, wider; condylus medialis subcircular in me- bpbm 165656. We follow Taylor (1998) for dial aspect; impresso ansae musculo iliofi- subspecies-level . bularis abuts suclus fibularis. Tibiotarsus: Measurements were taken with dial cali- impresso ligamentum collateralis medialis pers (Helios), rounded to the nearest 0.1 deep and wide; facies articularis femoris large; mm. To assess the degree of flightlessness in condylus medialis subcircular in medial as- each species, we performed a principal com- pect. Tarsometatarsus: corpus tarsometatarsi ponents analysis (PCA) using the software much wider than deep; medial sulcus hypo- package SPSS 13.0. Unweighted character tarsi not enclosed; fossa metatarsi I short and means (natural [base e] log-transformed) for shallow. Fossil Rails from Huahine . Kirchman and Steadman 285

Figure 3. Lateral (left) and dorsal (right) views of the skulls of Gallirallus. A, G. philippensis, uwbm 42866. B, G. storrsolsoni, holotype, bpbm 166036. C, G. owstoni, uf 39918. D, G. torquatus, ummz 228275. E, G. woodfordi, uf 39399. Scale bars ¼ 50 mm.

Gallirallus storrsolsoni Kirchman & Steadman, 168121 (l), 168150 (right [r]); radius, bpbm n. sp. 168056 (r); carpometacarpus, bpbm 168165 Figures 3B;4B;5B,E,H,K (l); synsacrum, bpbm 166020; femora, bpbm 168131 (r), dapt 27/105 (r); tibiotarsi, bpbm holotype. Complete cranium and ros- 166023 (l), 166032 (r), 168028 (l), 168046 trum, bpbm 166036 (Figure 3). (r), 168123 (r), 168149 (l), 168170 (l), dapt paratypes. Crania, bpbm 166021, 47 (r), 55 (l), 119 (l); tarsometatarsi, bpbm 166026, 168015, dapt 139; rostrum, dapt 166034 (r), 168124 (r), dapt 7 (r). 21; vertebrae, bpbm 166024, 166025, 166035, diagnosis. A medium-sized species of 168001, 168002, 168078, dapt 13, 14, 25, 60, Gallirallus (Table 1) distinguished from con- 61, 122, 143, 144, 145, 163; rib, bpbm 166018; geners in Oceania as follows. Skull (Figure sterna, bpbm 166017, 166027; humerus, bpbm 3): fossa temporalis deeply excavated, clearly 166022 (left [l]); ulnae, bpbm 166033 (l), emarginated by crista temporalis and extends TABLE 1 Skeletal Measurements (in mm) in Gallirallus, with Mean, Range, and Sample Size. F, female; M, male; U, sex unknown. Specimens of all available subspecies of G. australis and G. philippensis are combined, given that subspecific differences in size are much smaller than size differences between males and females.

Skeletal G. G. G. G. Element storrsolsoni G. owstoni G. australis ripleyi vekamatolu G. philippensis striatus G. torquatus G. woodfordi

Sex U M F M F U U M F M M F M F

Cranium length 33.7 33.3 31.6 42.9 40.0 — — 32.1 29.9 28.7 33.9 31.0 39.8 39.0 32.4–34.8 32.8–34.2 30.9–32.3 42.2–43.5 39.0–41.5 31.0–32.9 28.4–31.0 28.2–29.0 32.4–35.2 1 39.0–40.5 38.2–39.5 4 4 10 2 4 4 6 4 5 3 3 Rostrum length 38.8 40.7 37.1 52.0 46.0 — — 36.0 29.5 35.7 42.2 39.4 49.1 44.8 36.5–41.0 36.9–43.3 35.5–39.3 50.6–53.3 40.0–50.4 33.5–38.6 27.0–31.6 33.0–38.1 39.7–45.8 1 47.8–49.9 44.6–45.0 2 6 10 2 4 4 6 4 4 3 3 Sternal carina 9.1 10.7 10.4 12.0 10.6 — — 12.7 12.3 12.2 13.1 10.9 13.2 12.6 depth 1 9.7–11.6 9.9–11.4 11.5–12.5 9.9–12.0 12.0–13.5 11.6–13.5 11.6–13.0 12.2–14.1 1 12.6–13.8 12.2–13.4 7 10 2 4 6 6 3 5 3 3 Sternum width 14.6 11.7 11.3 20.9 19.2 — — 11.1 10.0 9.0 11.9 11.7 18.8 17.7 at coracoids 14.5–14.8 10.7–12.2 9.6–11.9 20.3–21.5 18.6–19.9 9.9–12.0 9.4–10.8 8.4–10.0 11.1–12.4 1 18.6–19.0 16.5–18.8 2 7 9 2 4 6 6 4 5 3 3 Humerus shaft 2.2 2.8 2.7 4.3 3.8 2.1 — 3.3 2.9 2.6 3.2 2.9 3.8 3.6 width 1 2.6–3.1 2.6–3.0 3.9–4.6 3.5–4.2 1 3.2–3.4 2.7–3.3 2.5–2.7 3.0–3.4 1 3.7–3.8 3.5–3.7 7 11 3 4 5 6 4 5 3 3 Ulna length 37.5 39.1 36.6 43.7 38.2 22.9 41.3 43.9 39.2 36.2 44.0 40.0 51.6 49.5 1 36.8–41.5 34.8–38.7 43.0–44.2 36.0–41.4 22.6–23.1 1 41.4–44.8 35.2–43.6 33.7–37.9 43.0–45.3 1 50.7–52.8 47.5–50.6 7 11 3 4 2 6 6 4 5 3 3 Ulna proximal 4.8 4.6 4.3 7.0 6.0 2.8 5.4 5.0 4.3 4.0 5.1 5.0 6.4 5.9 width 1 4.3–4.8 4.1–4.5 6.5–7.3 5.5–6.8 2.7–2.9 1 4.7–5.3 4.1–4.4 3.8–4.1 4.8–5.4 1 6.3–6.5 5.9–6.0 7 11 3 4 2 6 6 4 5 3 3 Femur length 49.4 56.0 52.4 80.6 69.9 39.8 — 53.8 47.3 45.1 55.5 50.6 72.1 69.6 1 54.0–59.2 49.2–54.6 78.7–82.3 66.1–73.4 1 51.3–54.9 45.0–49.7 42.2–46.9 53.3–57.5 1 72.0–72.3 69.0–70.3 7 11 3 4 6 6 4 5 3 3 Femur distal 9.2 9.5 8.7 16.5 14.3 6.7 10.9 8.7 7.4 45.1 9.3 8.5 13.3 12.7 width 1 8.9–9.9 8.4–9.0 15.9–17.6 13.2–15.0 1 10.7–11.0 8.4–9.1 7.0–7.9 42.2–46.9 8.6–9.9 1 13.2–13.4 12.5–12.9 7 11 3 4 2 6 6 4 5 3 3 Tibiotarsus 68.6 79.9 75.5 117.4 100.0 — — 76.9 66.7 62.1 84.2 74.6 105.0 102.8 length 1 76.1–84.6 72.9–79.8 116.6–118.7 93.4–105.0 71.7–80.3 63.0–70.6 58.2–64.3 80.9–87.5 1 100.6–108.8 99.0–105.8 7 11 3 4 6 6 4 5 3 3 Tibiotarsus 6.9 7.4 6.8 12.5 10.8 6.0 8.0 6.9 6.1 5.3 7.2 6.6 10.0 9.5 distal width 1 6.8–7.9 6.5–7.1 12.1–13.1 10.4–11.2 5.6–6.3 7.9–8.0 6.5–7.2 5.6–6.7 5.1–5.6 6.7–7.6 1 9.9–10.1 9.4–9.6 7 11 3 4 3 2 6 6 4 5 3 3 Tarsometatarsus 7.4 7.6 7.0 12.6 11.0 5.7 8.3 7.0 6.1 5.4 7.4 6.6 10.5 10.0 proximal 1 7.0–8.2 6.7–7.3 12.2–13.4 10.7–11.7 1 7.8–8.7 6.7–7.2 5.8–6.6 5.3–5.5 7.0–7.9 1 10.2–10.7 9.9–10.1 width 7 11 3 4 2 6 6 4 5 3 3 Tarsometatarsus 4.1 3.8 3.4 6.1 5.4 3.0 4.3 3.5 3.0 2.7 3.6 3.3 4.9 4.6 shaft width 1 3.5–4.0 3.2–3.8 5.6–6.4 5.2–5.6 2.8–3.1 4.0–4.6 3.1–3.6 2.8–3.3 2.5–2.9 3.3–3.8 1 4.8–5.0 4.4–4.7 7 11 3 4 2 2 6 6 4 5 3 3 Tarsometatarsus 2.8 2.8 2.5 4.8 4.1 2.2 3.2 2.8 2.5 2.1 2.7 2.6 4.0 3.8 shaft depth 1 2.5–3.0 2.4–2.8 4.5–5.0 3.8–4.6 2.1–2.3 3.0–3.3 2.6–3.0 2.3–2.8 2.0–2.2 2.2–2.9 1 3.9–4.2 3.7–4.0 7 11 3 4 2 2 6 6 4 5 3 3 Fossil Rails from Huahine . Kirchman and Steadman 287

Figure 4. Ventral (left) and lateral (right) views of the sterna of Gallirallus. A, G. owstoni, uf 39918. B, G. storrsolsoni, bpbm 166027. C, G. philippensis, uwbm 42866. Scale bars ¼ 50 mm. more caudally; cranium with prominent crista incisura intercondylaris wide, resulting from nuchalis transversae and a low, broad calveria an obtuse angle between the condylus medi- (as in G. woodfordi); in dorsal aspect, the pos- alis and condylus lateralis; juncture of condy- terior margins of the orbits abruptly angle lus medialis with facies caudalis of corpus away from the midline; the lamina parasphe- tibiotarsis abrupt rather than gradually slop- noidalis is well emarginated caudally. Ster- ing. Tarsometatarsus (Figure 5): corpus tar- num (Figure 4): spina externa of rostrum sometatarsi dorsoventrally flattened with a sterni absent. Humerus (Figure 5): incisura width-to-depth ratio (1.46) greater than in capitus narrows proximally in caudal aspect; other species (1.21–1.36); viewed medially, crista deltopectoralis rectangular, parallel to the proximal one-third of corpus tarsometa- corpus humeri; corpus humeri thin, round in tarsi slopes toward the hypotarsus rather cross-section. Ulna (Figure 5): corpus ulnaris than being perpendicular to facies dorsalis. straight and dorsoventrally flattened, more so etymology. Named after Storrs L. Ol- than even in flightless G. owstoni, G. vekama- son in recognition of his unparalleled con- tolu, or G. australis; impressio brachialis deep tributions to the , systematics, and and clearly emarginated. Synsacrum: broad paleontology of flightless rails on islands. in ventral aspect, gradually narrowing cau- remarks. Gallirallus storrsolsoni is a dally. Femur (Figure 8): corpus femoris ro- medium-sized species that, in overall size, bust, approaching but not surpassing the resembles G. owstoni, G. philippensis, G. stria- stoutness of G. vekamatolu. Tibiotarsus (Fig- tus, and G. torquatus. It is larger than G. ripleyi ure 5): proportionally short (as in G. australis); and smaller than G. australis, G. vekamatolu, Figure 5. Humeri (A–C), ulnae (D–F), carpometacarpi (G–I), tibiotarsi ( J–L), and tarsometatarsi (M–O)ofGalliral- lus philippensis (A, D, G, J, M, all from uf 39855), G. storrsolsoni (B, bpbm 166022; E, bpbm 166033; H, bpbm 168165; K, bpbm 166023; N, dapt 7), and G. owstoni (C, F, I, L, O, all from uf 42968). Scale bar ¼ 50 mm. Fossil Rails from Huahine . Kirchman and Steadman 289

TABLE 2 Principal Components Analysis Correlation Coefficients of 16 Skeletal Measurements from Eight Species of Gallirallus (See Text, Figure 6)

Correlation Coefficients

Skeletal Element PC1 PC2 PC3 PC4

Sternum, width at coracoids .265 .093 .045 .019 Sternum, depth .097 .177 .037 .027 Humerus, shaft width .219 .089 .077 .014 Ulna, total length .146 .110 .035 .003 Ulna, proximal width .217 .024 .013 .023 Ulna, shaft width .203 .049 .025 .030 Carpometacarpus, intermetacarpal space length .163 .165 .098 .022 Femur, total length .224 .021 .021 .002 Femur, distal width .293 .047 .004 .002 Femur, shaft width .255 .024 .002 .015 Tibiotarsus, total length .222 .014 .023 .031 Tibiotarsus, distal width .263 .048 .018 .011 Tibiotarsus, shaft width .263 .012 .002 .030 Tarsometatarsus, proximal width .258 .061 .004 .013 Tarsometatarsus, shaft width .253 .067 .029 .018 Tarsometatarsus, shaft depth .262 .037 .004 .004 Percentage total variance explained 85.81 10.65 2.33 0.61

and G. woodfordi. Gallirallus storrsolsoni has a ervation is unique; all other Gallirallus species greatly reduced carina sterni, small wing ele- described from fossils, G. ripleyi (, ments, and stout leg elements as in its flight- Cook Islands), G. huiatua (), and G. less congeners. The diagnosis of flightlessness vekamatolu (‘Eua, ), were considered is supported by morphometric comparisons to be flightless on the basis of limb-bone pro- with species of Gallirallus that are known to portions (Steadman 1987, Steadman et al. be either volant or flightless. Correlation 2000, Kirchman and Steadman 2005). coefficients of the first four principal com- ponents (PC), which together account for Genus Porphyrio Brisson, 1760 99.4% of morphological variance, indicate that PC 1 describes variation in overall size We refer three femora from the Fa‘ahia and the degree of reduction of the carina archaeological site (bpbm 166031, dapt 39, sterni, ulna, and carpometacarpus, and that 53) to Porphyrio rather than the other genera PC 2 is a description of keel and wing reduc- of large Pacific rails (Gallirallus, Gallinula, tion, and leg-bone robustness (Table 2). A Fulica) because of these characters: in proxi- plot of PC 1 versus PC 2, summarizing mal aspect, more obtuse angle formed at the 96.5% of morphometric variance, clusters G. junction of the impressiones obturatoriae storrsolsoni with flightless rather than volant and trochanter femoris; impressiones obtura- congeners (Figure 6). toriae more prominent, leading to a more The material from Fa‘ahia represents six concave proximoposterior area of corpus individuals, minimally. Because of the excel- femoris; similar size and position of the im- lent preservational environment at the Fa‘a- pressiones iliotrochanteria and linea inter- hia site, even the most delicate elements of muscularis caudalis; corpus femoris overall the skeleton of G. storrsolsoni are known, in- more slender; distal end of corpus femoris cluding one partial sternum (uf 166027) that not expanded laterally until the epicondylus still retains the anterior margin of the carina. lateralis is reached; rotular groove more nar- Outside the region, such pres- row; in posterior aspect, medial margin of 290 PACIFIC SCIENCE . April 2006

Figure 6. Plot of mean scores for eight species of Gallirallus on the first two principal components (summarizing 96.5% of variance) of 16 postcranial skeletal measurements. Hollow symbols are volant species; filled symbols are flightless species. The line indicates a hypothesized threshold for flightlessness. the condylus medialis oriented roughly paral- farther (more distad) along corpus femoris lel to the shaft rather than diagonal. than in P. paepae; in lateral aspect, corpus femoris straighter than in all but P. paepae; Porphyrio mcnabi Kirchman & Steadman, n. in lateral aspect, crista trochanteris more sp. rounded (less flared) on dorsal surface than Figures 7A,8D in P. paepae; most proximal section of linea intermuscularis caudalis (¼ dorsalis) weakly holotype. Nearly complete right femur, developed (thicker and more protrudent in bpbm 166031 (Figures 7, 8). P. paepae). paratypes. Distal left femur, dapt 53. etymology. Named after Brian K. Left femur lacking distal end, dapt 39. McNab in recognition of his important re- diagnosis. A small species of Porphyrio search on the evolution and physiological (Table 3) distinguished from congeners as ecology of flightless birds, especially rails, on follows: impressiones iliotrochanteris extends oceanic islands.

—————————————————————————————————————————————————f Figure 7. Cranial (upper) and caudal (lower) views of femora of Porphyrio. A, P. mcnabi, holotype, bpbm 166031. B, P. paepae, bpbm 165649. C, P. martinicus, uf 42419. D, P. porphyrio, uf 39407. Scale bars ¼ 50 mm.

Figure 8. Femora of Gallirallus and Porphyrio in cranial aspect. A, G. philippensis, uf 39855. B, G. storrsolsoni, dapt 27/105. C, P. martinicus, uf 42419. D, P. mcnabi, bpbm 166031. Scale bar ¼ 50 mm.

TABLE 3 Selected Skeletal Measurements (in mm) of the Femur in Porphyrio with Mean, Range, and Sample Size. F, female; M, male; U, sex unknown.

P. P. P. porphyrio P. [porphyrio] P. [porphyrio] P. P. Femur mcnabi paepae samoensis pulverulentus poliocephalus martinicus alleni

Sex F? U M F M F M M F M F

Total 49.4 52.1 81.3 70.0 78.2 75.4 68.8 53.8 50.7 46.1 44.7 length 1 51.7–52.5 76.4–86.5 64.7–72.8 77.9–78.4 1 1 52.0–55.4 47.7–52.4 1 1 2 3 3 2 5 6 Shaft 3.9 4.1 5.9 5.0 5.2 5.1 4.8 3.6 3.4 3.0 2.9 width 1 3.9–4.2 5.6–6.1 4.5–5.4 5.1–5.3 1 1 3.3–3.7 3.2–3.6 1 1 3 3 3 2 6 6 Distal 9.2 9.5 13.7 12.4 13.8 13.4 11.7 8.5 7.9 7.0 6.4 width 1 9.5–9.5 13.0–14.8 10.9–13.8 13.4–14.2 1 1 8.1–8.7 7.4–8.8 1 1 2 3 2 2 6 6

Note: Some measurements of P. porphyrio and P. martinicus are from Steadman (1988). Fossil Rails from Huahine . Kirchman and Steadman 293

remarks. Porphyrio mcnabi is a small spe- (McNab 1994a, 2002), suggests that selection cies that, in overall body size, resembles P. for reduced pectoral and wing musculature is paepae and P. martinicus. It is larger than P. the likely cause of insular avian flightlessness. flavirostris and P. alleni and is exceeded in Skeletons of flightless rails are distin- size by P. porphyrio, P. mantelli, P. hochstetteri, guished from those of volant relatives by hav- P. kukwiedii, and Porphyrio undescribed spe- ing reduced sternal keels (carina sterni) and cies A and B (see later in this section). The shorter, thinner wing bones. Livezey (1998, three femora of P. mcnabi were among the 2003) has shown that the degree of reduc- 50 bones listed as ‘‘Gallirallus new sp.’’ in tion of pectoral elements varies greatly even Steadman and Pahlavan (1992). It is likely among flightless species. Indeed there appears that bpbm 166031 represents an adult female to be a continuum of wing reduction among (smaller), whereas dapt 39 (left) and dapt flightless rails that mirrors the graded reduc- 53 (right) represent a single juvenile male tion in energy expenditure (McNab 2002). (larger). That dapt 39 and 52 are from a ju- Our morphometric analysis of Gallirallus venile is supported by their porous surfaces, skeletons indicates that some species, such as thin-walled shafts, and being slightly more G. australis, have greatly reduced carina sterni gracile than would be expected in an adult. and wing elements, whereas other extant spe- Lacking elements of the wing or pecto- cies known to be flightless, such as G. owstoni, ral girdle, we cannot say whether Porphyrio seem to be near the threshold of flight. mcnabi was flightless. The extant P. martini- cus, P. flavirostris, P. alleni (all small), and P. Biogeography porphyrio (large) are all volant. The four very large, extinct species (P. kukwiedii of New Of the G18 living and extinct species of Gal- Caledonia, P. mantelli of New Zealand, Por- lirallus sensu lato that have been named, all phyrio undescribed species A of New Ireland, but two are flightless species endemic to Oce- Bismarck Archipelago, and Porphyrio unde- ania on single islands or on multiple islands scribed species B of Buka, Solomon Islands) that were connected during the late Pleisto- all were flightless, as is the very large, extant cene period of lowered sea levels (Steadman P. hochstetteri of New Zealand (Balouet and 1987, in press, Diamond 1991, Mayr and Di- Olson 1989, Worthy and Holdaway 2002, amond 2001). The two extant, volant species, Steadman in press). The small, extinct P. pae- G. torquatus and G. philippensis, are sympatric pae of the Marquesas had somewhat reduced in the Philippines, Sulawesi, and New Guinea, wings but perhaps was still volant (Steadman although the latter species is very widespread, 1988). its distribution extending south to and New Zealand and east in Oceania to discussion . Flightless species of Gallirallus have evolved on nearly all major archipelagos in Flightlessness Oceania from the Ryukyu Islands of southern Rails are developmentally predisposed to Japan, south to New Zealand’s Chatham become flightless (Olson 1973b, Feduccia Islands, and east to the Society Islands. Bones 2000), and scores of neotenic flightless spe- (still undescribed) of Gallirallus also have cies have evolved independently on oceanic been found in archaeological sites on four islands that lack indigenous placental carni- islands in the Marquesas (Steadman 1989a, vores. The hypothesis that flightlessness Steadman and Rolett 1996). Gallirallus ap- evolves as a means of energy conservation is parently never made it as far northeast in supported by the basal metabolic rates in Oceania as the or as far flightless species of insular rails being lower southeast in Polynesia as Henderson Island, than those of their volant relatives (McNab both of which have good Holocene fossil 1994a,b). This finding, coupled with the - bird records ( James and Olson 1991, Olson servation that metabolic rate also correlates and James 1991, Wragg 1995, Steadman in positively with relative pectoral muscle mass press). 294 PACIFIC SCIENCE . April 2006

Gallirallus storrsolsoni is not the only species southern and eastward across of flightless rail known from the Society Is- southern , Indonesia, and Australia to lands. The extinct G. pacificus was discovered Oceania as as Fiji, Tonga, Samoa, on Tahiti by naturalists from Captain James and Niue (Taylor 1998:464, 465). No species Cook’s second voyage (1777), but no speci- of Porphyrio inhabit East Polynesia today. mens exist, and the species is known only The radiation of certainly or presumably from a painting by . The plum- flightless species of Porphyrio is confined to age, soft-part colors, and bill shape make it Oceania (Balouet and Olson 1989, Worthy clear that pacificus is correctly accomodated and Holdaway 2002, Steadman in press). in Gallirallus. Measurements made by Storrs The only flightless species that still exists is L. Olson in 1998 (pers. comm.) from For- P. hochstetteri from South Island, New Zea- ster’s original, full-scale painting in the Brit- land. Known extinct forms of Porphyrio are ish Museum of Natural History indicate that P. mantelli (North Island, New Zealand), P. G. pacificus was a much smaller rail than G. albus (Lord Howe Island), P. kukwiedii (New storrsolsoni. For example, the culmen (with Caledonia), and Porphyrio undescribed sp. A epidermal sheath) in G. pacificus is 28.8 mm, (New Ireland) and sp. B (Buka, Solomon Is- whereas the rostrum (without epidermal lands). Each of these species was as large as sheath) in G. storrsolsoni is 36.5–41.0 mm or larger than the massive P. hochstetteri, (Table 1). The tarsus (incuding scutes) in G. which is the largest extant form. pacificus is 33.8 mm, whereas the tarsometa- Finally, Porphyrio paepae was a smaller, tarsus (without epidermal sheath) in G. storrs- probably flightless that is known olsoni is ca. 48.5 mm, based on a composite from and , two islands only of two incomplete specimens (dapt 7, bpbm 3 km from each other in the Marquesas 166034). Gallirallus pacificus is presumed to (Steadman 1988). At the time of its descrip- be flightless on the basis of the short wings tion, P. paepae was the only species of Por- in the painting; although this is likely, in the phyrio known from East Polynesia. The absence of specimens it cannot be verified in discovery of P. mcnabi in the Society Islands the painting. The possible former existence helps to bridge the formerly huge distribu- of G. pacificus on Mehetia (Taylor 1998), a tional gap of the genus (from Niue to the small island 110 km east-southeast of Tahiti, Marquesas) and strengthens the likelihood is unsubstantiated and doubtful. that a substantial radiation of swamphens Based on its geological and geographical once existed in East Polynesia. At this point, setting, any flightless species on Huahine is we cannot say whether this radiation was of likely to have been endemic to the island. or origin. Osteologi- The geological age of Huahine is ca. 3 mil- cal synapomorphies that ally P. paepae and P. lion yr (Dickinson 1998). It is an eroded vol- mcnabi with either the New World species (P. canic island surrounded by a broad fringing martinicus, P. flavirostris) or the Old World reef, outside of which the water becomes P. porphyrio s.l. have not been discerned. very deep. The nearest island is Ra‘iatea, ca. Given the propensity in P. martinicus and P. 35 km to the west. Huahine never was con- flavirostris for unpredictable, long-distance nected to any other island, even during the dispersal (Remsen and Parker 1990), the geo- lowered sea levels of Pleistocene glacial inter- graphic origin of East Polynesian Porphyrio vals, when Ra‘iatea still would have been ca. species remains an open question. 30 km away. The genus Porphyrio is widespread in trop- Avian Extinction on Huahine ical and subtropical lowlands. Two extant, volant species, P. martinicus and P. flavirostris, The Fa‘ahia archaeological site contained the are confined to the . All other spe- bones of 15 resident species of , three cies are from the Old World, with extant, vo- migratory species of shorebirds, and 16 spe- lant P. alleni in Africa and the larger, extant, cies of resident land birds (Steadman and volant P. porphyrio very widespread from Pahlavan 1992, Steadman in press). Of the Fossil Rails from Huahine . Kirchman and Steadman 295

15 seabirds, 12 no longer occur on Huahine, his measurements of Forster’s painting of including the extinct gull Larus utunui (Stead- Gallirallus pacificus. We thank Storrs Olson man 2002). The other 11 species of seabirds and an anonymous reviewer for insightful still are found, at least locally, elsewhere in comments that improved the manuscript. Oceania. Each of the migratory shorebirds, on the other hand, may visit Huahine season- Literature Cited ally today. Of the 16 species of land birds from Fa‘ahia, 13 no longer occur on Hua- Balouet, J. C., and S. L. Olson. 1989. Fossil hine. They consist of the locally extripated birds from late Quaternary deposits in heron Butorides (Ardeola) striatus, rail Porzana New Caledonia. Smithson. Contrib. Zool. tabuensis, dove Gallicolumba erythroptera, pi- 469. geons Ducula galeata and D. aurorae, and war- Baumel, J. L., and L. M. Witmer. 1993. Os- bler Acrocephalus caffer. Aside from Gallirallus teologia. Pages 45–132 in J. J. Baumel, storrsolsoni and Porphyrio mcnabi, the extinct A. S. King, J. E. Breazile, H. E. Evans, and species of land birds from Fa‘ahia include J. C. Vanden Berge, eds. Handbook of the doves Gallicolumba nui and Macropygia avian : Nomina anatomica avium, arevarevauupa, lorikeets Vini vidivici and V. si- 2nd ed. Nuttall Ornithological Club, notoi, and starling Aplonis diluvialis (Steadman Cambridge, Massachusetts. and Zarriello 1987, Steadman 1989b, 1992). Conte, E., and A. Anderson. 2003. Radiocar- The survivors are the very widespread bon ages for two sites on , Mar- heron Egretta sacra and two species endemic quesas. Asian Perspect. 42:155–160. to the Society Islands, the dove Ptilinopus pur- Diamond, J. M. 1991. A new species of rail puratus and kingfisher Halcyon tuta. Also ex- from the Solomon Islands and convergent tant on Huahine is the very widespread evolution of insular flightlessness. Auk Anas superciliosa, which was not found among 108:461–470. the bird bones from Fa‘ahia. Two additional Dickinson, W. R. 1998. Geomorphology and species of land birds, the lorikeet Vini peruvi- geodynamics of the Cook–Austral Island ana and swiftlet Collocalia leucophaea, are un- Seamount Chain in the South Pacific known at the Fa‘ahia site but were recorded Ocean: Implications for hotspots and from Huahine in the nineteenth century, al- plumes. Int. Geol. Rev. 40:1039–1075. though they no longer persist on the island Dye, T., and D. W. Steadman. 1990. Polyne- (Steadman in press). Altogether, 19 species sian ancestors and their world. Am. of land birds have been recorded from Hua- Sci. 78:209–217. hine, which is the same number that is known Feduccia, A. 2000. The origin and evolution from much-larger Tahiti, which has no fossil of birds, 2nd ed. Yale University Press, record of birds. If the prehistoric bird com- New Haven, Connecticut. munity of Huahine were entirely revealed, James, H. F., and S. L. Olson. 1991. Descrip- we believe that it also would include a species tion of thirty-two new species of birds of Prosobonia sandpiper, Cyanoramphus parrot, from the Hawaiian Islands: Part II. Passer- and Pomarea monarch, three genera recorded iformes. Ornithol. Monogr. 46. on at least two other islands in the Society Kirch, P. V. 1984. The evolution of the Poly- Group. nesian chiefdoms. Cambridge University Press, Cambridge, United Kingdom. acknowledgments ———. 1986. Rethinking East Polynesian . J. Polynesian Soc. 95:9–40. For access to modern specimens, we thank ———. 2000. On the road of the winds: An the curatorial staffs of the eight museums archaeological history of the Pacific islands listed in Materials and Methods. Carla Kishi- before European contact. University of nami and Yosihiko Sinoto kindly provided ac- California Press, Berkeley. cess to the prehistoric bones from Huahine. Kirch, P. V., and J. Ellison. 1994. Palaeoen- We are grateful to Storrs Olson for sharing vironmental evidence for human coloniza- 296 PACIFIC SCIENCE . April 2006

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